lepidopteranLepidopteraany of more than 155,000 species of butterflies, moths, and skippers. This order of insects is second in size only to Coleoptera, the beetles.

Because of their day-flying habits and bright colours, the butterflies are more familiar than the chiefly night-flying and dull-coloured moths, but the latter are far more varied and abundant. The skippers are a worldwide group intermediate between butterflies and moths. With the exception of a few moths, all adult lepidopterans have two pairs of wings. The name Lepidoptera is derived from the Greek, meaning “scaly winged,” and refers to the characteristic covering of microscopic dustlike scales on the wings.

The life cycle of lepidopterans consists of four stages: egg, larva (caterpillar), pupa (chrysalis), and adult (imago). The larvae do most of the eating, with the majority feeding on foliage, although many species eat stems, roots, fruits, or flowers. A number of moth and a few butterfly larvae are serious pests in agriculture and forestry. The adults of many species are important for their role in pollination, which occurs as they visit flowers for nectar. In most land environments the lepidopterans are ecologically important because they transform large amounts of plant matter into animal matter and in turn serve as food for many other groups of animals.

Many members of the order, especially the butterflies, have appealed to the human imagination for thousands of years as symbols of fragile and ephemeral beauty. References to them abound in literature, and they have been depicted in many paintings, have inspired the designs of jewelry, ornaments, and textiles, and have even occurred in many heraldic devices and on postage stamps.

Size range and distribution

Moths, butterflies, and skippers show great diversity in size and development rates. Some moths have wingspans as small as 4 mm (0.13 inch), whereas the largest moths and butterflies measure nearly 30 cm (about 1 foot). Fast-developing species may complete their development in as little as three weeks, while slower ones may require as long as two or even three years.

Lepidopterans live on every continent except Antarctica. Though they are far more numerous and diversified in the tropics, some species survive at the limits of polar vegetation. There are many successful species in nearly every environment, from arid deserts and high mountaintops to marshes and tropical rainforests. Most have become adapted for life in relatively restricted ecological niches and may be limited to only one or a small group of food plants, often to only a single part of one plant. They are therefore seldom abundant in more than one type of habitat. However, a few species have broader food habits and may occasionally reach peaks of abnormal abundance and defoliate large areas of deciduous forest or grassland. Some of the insects most injurious to human interests in agriculture and forestry are members of the Lepidoptera.

The large family-level groups and many smaller ones are worldwide in distribution, with owlet moths (family Noctuidae), measuring worm moths (family Geometridae), pyralid, or snout, moths (family Pyralidae), brush-footed butterflies (family Nymphalidae), and skippers (family Hesperiidae) being dominant elements of the insect fauna everywhere. A few families are characteristically more abundant in one faunal region than in others. Central and South America are characterized by great diversity of American false tiger moths (family Notodonidae) and ctenuchid moths (family Arctiidae). The North American (Nearctic) and Eurasian (Palaearctic) families show much evidence of close connections, chiefly between Asia and western North America. Each region has, however, many distinctive genera.

Many lepidopterans exist only in isolated colonies as relict (remnant) populations, cut off from relatives elsewhere by geologic or climatic changes. Australia and New Zealand have unusually diverse relict populations of the primitive mandibulate moths (family Micropterigidae) and swifts, or ghost moths (family Hepialidae). In North America, Europe, and Asia, many relict species have survived since the last ice age on isolated southern mountaintops.

Importance

Many hundreds of Lepidoptera injure plants useful to humans, including some of the most important sources of food, fabrics, fodder, and timber. The great majority of the injurious species are moths, and the detrimental life stage is always the larva. However, unlike members of other insect orders, lepidopterans do not act as carriers of plant diseases, nor are any of them parasites of or injurious to humans. However, some species feed on open wounds or bodily secretions of wild or domestic animals.

The list of valuable plants subject to damage by lepidopterans is a long one, including many grains, sugar beets and sugarcane, cotton, tobacco, some root crops and leaf crops, many fruits, and timber and shade trees. The damage may involve the leaves, stems, roots, or fruit. Woolens, furs, silk, and even feathers are eaten by fungus moths (see tineid moths) of several genera (clothes moths). The wax moth (Galleria mellonella) causes considerable damage in beehives.

A few Lepidoptera are directly beneficial to humans. Nearly all silk is obtained from the domesticated silkworm (Bombyx mori), which is originally from China. Other silks such as shantung and tussah are the products of various Asiatic giant silkworm moths (family Saturniidae). The larvae and sometimes the adults of a few species are used for food. The larvae of one skipper (Rhopalocampta libeon) are collected in large quantities in the Congo, and the 10-cm (4-inch) caterpillars of giant skippers (family Megathymidae), known in Mexico as gusanos de magüey, are both consumed domestically and canned and exported for consumption as hors d’oeuvres. The South American cactus moth (Cactoblastis cactorum) has been highly beneficial in weed control, clearing more than 150 million ha (60 million acres) in Australia of alien prickly pear cactus. Doubtless, humans also benefit from much unrecognized weed eating by caterpillars and flower pollination by adults.

Many lepidopterans are valuable in biological research, including work in ecology, biogeography, systematics, genetics, and physiology. Much of the present knowledge of endocrine controls of insect development has come from studies of the silkworm moth and its relatives. The study of the British peppered moth (Biston betularia) has profoundly influenced ideas about rates of evolutionary change. An increase in the proportion of dark moths, a change thought to be brought on by airborne soot produced during the Industrial Revolution, has been termed industrial melanism.

Natural history

Feeding habits

The habits of Lepidoptera are extremely diverse, depending on the adaptations of the species or group to climate, environment, type of food plant, way of feeding, and many other factors. The great majority of food plants are conifers and flowering plants, but primitive plants such as mosses, liverworts and ferns, and some lichens are eaten by a few groups. Nearly all parts of the plant are eaten by various specially adapted caterpillars. Flowers are eaten by many larvae, including plume moths (family Pterophoridae), with nectar being consumed by many adults. Cones and fruits and their seeds are eaten by others, such as yucca moths (family Incurvariidae), leaf roller moths (family Tortricidae), and some owlet moths. Some seed eaters such as the flour moth (genus Ephestia) have become household pests, feeding on stored grains and cereals. Buds or soft, succulent stems are bored into by members of many families. Several lepidopteran groups—for example, the pine moth (Rhyacionia)—specialize on the terminal shoots of conifers. Various groups feed on the turf of grasses and sedges. The carpenter (family Cossidae), ghost (family Hepialidae), and clearwing moths (family Sesiidae) bore into woody stems and rootstocks. Carpenter moths in particular tunnel deep into hardwoods. A good many lepidopterans, especially fungus moths (family Tineidae), scavenger moths (family Blastobasidae), and snout moths (family Pyralidae), feed on dead and decaying plant matter, particularly moldy debris. In comparison with other orders of insects, relatively few Lepidoptera live in plant galls or eat animal matter. Several families contain leaf miners, especially Gracillariidae (leaf miner moths), Elachistidae (grass miner moths), and Tischeriidae (trumpet leaf miner moths).

Life cycle

Egg

The number of eggs laid varies greatly from fewer than a hundred in some species to more than a thousand in others. The eggs are almost always laid in a specific way, usually on or in a suitable food plant. Eggs of many species are laid singly and widely dispersed; in others they are laid in masses that may be covered with a hardened secretion from the female’s abdominal glands. In a few groups the terminal segments of the female’s abdomen are greatly elongated and bladelike, and the eggs are laid in soft plant tissues or in narrow slits or crevices. In some of the primitive moths (family Prototheoridae) and in some butterflies, the female may merely scatter the eggs in flight in the general vicinity of a suitable food plant. Development of the embryo and emergence of the young larva is often controlled by a mechanism of physiologically enforced inactivity (diapause), which has the effect of timing the emergence of the larva to coincide with suitable conditions of weather and the growth of the food plant. Respiration in the egg is carried on through an aeropyle, a system of air passages in the shell (chorion) that enables oxygen exchange with the environment to occur whether the egg is dry or wet. In a few species of scavenger moths and pierid butterflies (see white and sulfur butterflies), the larvae hatch in the uterus of the female.

Larva, or caterpillar

The larva is the chief, and often the only, feeding stage of the life cycle. Its function is simply to transform very large quantities of plant matter into animal matter and to stay alive during the process. Most larvae molt four or five times as they grow, shedding their exoskeleton in a process called ecdysis. With plant matter being relatively easy to find and eat, it is not surprising that moth and butterfly larvae are fundamentally quite uniform, despite their apparent diversity.

Most larvae continually spin silk threads that adhere to the surfaces on which they live, thus giving them an almost unshakable grip. Silk is also used in the construction of various nests, cases, and shelters that protect the larvae from the elements and from enemies. In moths, silk also serves in the final cocoon, in which the larva will change into a pupa. The larvae of leaf miners are well adapted for life inside a flat leaf, as they are very small, greatly flattened, and more or less legless. The larvae of borers are relatively plain and unornamented, while larvae that live in the open, exposed to the attacks of predators, have evolved a multiplicity of protective adaptations of form, colour, texture, behaviour, and chemical repellents. The proper timing of larval development and activities is important for species in which a dormant period is necessary. For example, in admiral butterflies (Limenitis) the larvae of the summer generation complete development without pause, but those that hatch in the fall hibernate while still very young. In certain groups larval development is very slow (for reasons not understood), so that there may be only one generation per year. The larvae of the carpenter moths that feed on nutrient-poor wood may take two years to develop. Larvae of many Arctic and northern species regularly have two-year cycles, because of both cooler temperatures, which slow growth rates, and the shorter growing season of their food plants. The usual number of molts (ecdyses) is four or five, but some of the small leaf miners molt only twice. When starved, the larvae of clothes moths (Tineola) have been known to have a dozen molts, sometimes accompanied by a decrease in size.

Larvae seldom travel far from the plants where they begin life. However, in some species there is dispersal of very young larvae, which hang on silk threads and are blown by the wind. Swarms of armyworms (Pseudaletia) may travel long distances along the ground, driven by crowding and lack of food. Just before pupation many larvae stop eating and crawl some distance before settling down to pupate.

Pupa, or chrysalis

The larval stage is followed by the pupa, a resting stage in which the caterpillar undergoes a major rebuilding of body tissues to emerge as a mature adult. For moths, many species pupate in the soil, with little or no cocoon; many others form cocoons in the soil, in leaf litter, or under loose bark. Some cocoons are fastened to twigs or branches or rolled in leaves. The cocoons of leaf miners are usually formed in the “mine” or near it. Cocoons are commonly of silk alone but may also incorporate larval hairs, secretions or waste products, or chewed wood pulp, trash, or leaves. There may be a special seam on the cocoon to facilitate the emergence of the adult moth. Skipper larvae make a flimsy cocoon, generally in a curled leaf where it has been feeding. Of the true butterflies, only the satyr and parnassian butterflies make cocoons; all others pupate naked. In many species the pupa hangs in a head-down position from a silk pad by a stalk (cremaster). The chrysalis of some sulfur butterflies (family Pieridae), swallowtails (family Papilionidae), and gossamer-winged butterflies (family Lycaenidae), is supported in a head-up position by a threadlike silk girdle about the body.

The duration of the pupal stage differs greatly within various groups and sometimes even in different generations of a single species. In small, rapidly developing species, it may last only a week or 10 days during the summer or for many months in cases of hibernation or estivation. Pupae have been known to remain alive for three years in abnormal conditions and still produce adults. The danger of desiccation is greatest in small and exarate pupae (those in which the appendages are not fixed to the body by a skin or sheath) and least in large or compact ones.

The adult

Although fully formed, the adult may remain quiescent within the pupal case for a long time until conditions are right for its emergence. To escape from the cocoon, the pupae of some groups have cocoon-cutting structures such as the movable mandibles of certain primitive moths, the saw-toothed structures on the head of blotch leaf miners, and the bladelike structures on the wing bases of giant silkworm moths. In puss moths (Cerura) and some others, the cocoon is partially dissolved by alkaline secretions.

Once out of the cocoon, the adult crawls upward to where it can hang with the head up and the back down. The insect then forces its body fluids into the thorax by contracting its abdomen and pumping blood into the unexpanded wing pads, causing them to take on the size and shape of the adult wings. The adult may be able to fly in a few minutes or may have to hang for several hours until its wings have stiffened sufficiently for flight.

Whereas the larva is the nutritive stage of the life cycle, the adult is the reproductive stage. Its mobility is necessary to bring males and females together for copulation and to disperse the species into new areas. Nutrition is also an essential adult function in many primitive groups, such as the mandibulate moths (family Micropterigidae). In most of the highly mobile species, much of the energy necessary for flight is obtained from nectar or other liquid foods taken in by the adult. In a few groups the adult mouthparts have become so reduced that they do not function. In these species, all nutrition is obtained during the larval stage.

Behaviour

Food selection by the adult

Adults locate their food sources by both sight and scent, the former being especially important in diurnal species and the latter in many diurnal and in most nocturnal species. The chief source is floral nectar, but sap (especially if fermenting), overripe fruits, homopteran honeydew (sugar-containing secretions from homopteran insects), fecal matter, and carrion are sometimes used. There are mutualistic relationships of a broad sort between species of lepidopterans with flower-visiting adults and the plants whose flowers they visit and pollinate. However, these relationships are seldom specific or obligate, since only rarely are the plant and the lepidopteran mutually dependent. Exceptions exist among some orchids and members of the morning glory family, both of which have very deep tubular flowers. These appear to be pollinated only by certain hawk moths (family Sphingidae) with very long “tongues” (proboscises). The mutualism of the yucca moths and yucca plants is obligate in that the moth larvae feed only in yucca fruits and the latter can develop only from moth-pollinated flowers. The female yucca moth has special tentacles on the mouthparts, with which it gathers and carries balls of pollen (see community ecology: The coevolutionary process).

Courtship and mating

The sequence of mating activities is usually initiated by the female, which gives off specific odorous substances (pheromones) that attract males, sometimes even before she emerges from the pupa. These are detected by structures (scent sensilla) on the male’s antennae. Males with very large, feathery antennae, such as those of the giant silkworm moths, can locate females from 5 to 6 km (3 to 4 miles) away and may form courting swarms about them. A species may have a particular time of day or night for this release of pheromones. Males of some species, such as the European ghost moth (Hepialus humuli), at dusk may form dancing swarms into which the females fly.

Once the male has located the female, a more or less elaborate courtship may ensue. The sight of characteristic colours and patterns may be a requisite for this among the brightly coloured butterflies. Male pheromones also may play an essential part. Distributed from special scent scales (androconia) on the wings, body, or legs, the pheromones ensure the receptivity of the female. Finally, the accessory genitalic structures must fit together, not only mechanically but also in such a way that stimulates sensory nerve organs of the female. Some butterfly courtships are very active. In the sulphurs, for example, the male and female fly high in the air, with elaborate behavioral sequences in which each act serves as a stimulus for the next act. Such highly specific sequences are important barriers to hybridization between otherwise similar species.

The males of many species show definite territorial behaviour, defending a particular perch and area against the intrusion of other males but darting out at passing females. Among butterflies, certain admirals (Limenitis), coppers (Lycaena), and hackberry butterflies (Asterocampa) are noted for this behaviour. Congregations of males of many butterflies about hilltops may have a certain territorial and premating function. So may the large aggregations of various butterflies living in wet places.

Migration

Many Lepidoptera are famous migrators. The American monarch butterfly (Danaus plexippus) is the only species known to perform an annual long-distance two-way migration; i.e., the same individuals fly southward in the autumn and northward in the spring. Monarchs have also crossed the Pacific Ocean, colonizing Hawaii and Australia; occasionally they reach Africa and Europe. The cosmopolitan painted lady (Vanessa cardui) stages mass flights nearly everywhere it is found. However, these are one-way flights and are therefore emigrations rather than true migrations. Many other species in Europe and North America fly northward in large numbers, often reaching regions in which they cannot survive the winters. In Europe the painted lady and many moths reach Britain and Scandinavia from central Europe. In North America the painted lady, cloudless sulphur (Phoebis sennae), and many owlet moths often reach Canada. Many spectacular emigrations occur in the tropics, where swarms numbering in the millions may fly out to sea and become lost. The best-known group having these mass movements are pierid butterflies, but mass flights of certain large day-flying swallowtail moths (Urania leila and U. fulgens) have also been recorded. The usual explanation of such mass population movements, that they serve to extend the range of the species, is far from adequate in many instances.

Studies employing radar technologies revealed that migratory lepidopterans can travel at speeds of 100 km (62 miles) per hour by selecting fast-blowing wind currents a few hundred metres above ground. They also are able to identify currents that will carry them in the direction of their destination and can fine-tune their direction in response to crosswinds, enabling them to maintain their course. The deliberate selection of favourable winds combined with corrections for wind drift have been estimated to extend by 40 percent the distance that the insects are able to migrate. These behaviours, which are fundamental to the success of long-distance lepidopteran migrations, parallel those observed in migratory birds.

Ecology

Environmental hazards

As primary consumers of green plants, lepidopterans are enormously important in food chains, not only because of the very large number of species in the order and the diversity of their food habits but also because of their abundance. Lepidopterans, in turn, are eaten by a host of predators, parasites, and scavengers. All stages in their life cycles are under continual attack.

The major invertebrate predators on lepidopterans include centipedes, spiders, mantids, bugs (homopterans), ground beetles, ants, and both social and solitary wasps. Important predators among vertebrates include toads and tree frogs, lizards, birds, rodents, bats, and monkeys. The invertebrates generally locate their prey by scent or sight, whereas most of the vertebrates hunt by sight. The exception are the bats, which hunt by acoustic echolocation (the so-called bat “sonar”).

The chief groups of parasites that attack lepidopterans are tachinid flies and many wasps, chiefly the ichneumon, chalcid, and cynipid wasps. More precisely called parasitoids, these insects probably have a greater impact on caterpillar populations than do the direct predators. Female parasitoids locate suitable hosts, chiefly by scent, and lay their eggs in, on, or near them. The parasitoid larvae live inside their hosts, gradually feeding on their tissues and almost invariably consuming them almost completely. Unless some of the caterpillars’ toxic or repellent secretions serve to discourage them, lepidopterans seem to have evolved few defenses against parasitoids. The high reproductive rate of lepidopterans is important in countering losses to parasitoids as well as other adversities.

Small red chigger mites often ride about on adult lepidopterans but probably do them no harm. However, a few other mites live and breed in the tympanic cavities of owlet moths, destroying their auditory structures. Curiously, these mites regularly settle in only one of a moth’s two tympanic cavities and thus only half-deafen it. It is believed that by leaving the moth with one good “ear,” the mite reduces the likelihood of the moth, and hence of the mite itself, being captured and eaten by a bat. Lepidopterans are also subject to attack by a considerable number of protozoa, roundworms, bacteria, viruses, and fungi that affect the larvae chiefly during peaks of abundance and crowding. Some of these organisms have been used by humans as a means of controlling injurious species.

Protection against danger

It is chiefly against the visually hunting predators that lepidopterans have evolved a multiplicity of defense mechanisms. The adults of many groups, such as skippers, many butterflies, hawk moths, and many underwing moths, have fast and erratic flight. When escaping, they dart or fall to cover and often remain immobile for some time. They have a good chance of survival, especially if their colour matches their surroundings. Larvae, especially when small, drop suddenly when disturbed, either dangling from a silk thread or falling to concealment on the ground. The larvae of some owlet moths can jump several inches. Dense, loose hairs and scales make many moths slippery and may facilitate their escape from sticky spiderwebs.

Certain owlet moths and possibly some measuring worm and snout moths that are subject to predation by bats are able to receive and identify the bats’ navigational sound pulses. Upon hearing bat pulses, these moths perform radical evasive maneuvers when the bat sound is loud (hence close) or dive to the ground when the bat pulse is weaker (indicating that the danger is farther away).

Targets such as prominent coloured eyespots or tails on the hind wings attract attention and focus the attacks of predators onto parts of the body that are less vulnerable to injury. Such spots are likely to be seized and torn off, but this does the moth or butterfly no real harm and gives it time to escape without vital injury.

Many species manage to hide very effectively from predators. Cutworms and other caterpillars hide in litter by day and feed only at night. Many moths hide in crevices or under loose bark. Some of these seem to have especially flat bodies for this purpose. Hibernating butterflies spend the winter in hollow trees or among dead leaves, where they hang immobile. The larvae of a great many moths, most skippers, and many butterflies live in individual nests of rolled, folded, or webbed leaves or grass. Sod webworms live in silk-lined tunnels in turf. Wood borers, especially those in rootstocks and deep tunnels, are relatively secure. Many larvae aggregate in communal nests—e.g., tent caterpillar moths (Malacosoma), the ermine moth (Yponomeuta), and the Mexican social white butterfly (Eucheira socialis). The larvae of the bagworm moths (family Psychidae) and casebearer moths (family Coleophoridae) live and pupate in individual portable cases that are often masked with bits of leaf or twig. Some larvae, such as those of the green measuring worm moths (Synchlora), attach bits of leaves or flower petals to themselves. Cocoons are frequently camouflaged with leaves or debris. The chrysalis of puss moths (Cerura) and some dagger moths (Apatela) is hard, woody, and inconspicuous.

Camouflage is a form of concealing coloration. Such a cryptic, or hidden, appearance occurs when the natural coloration or pattern of an organism allows it to blend in with its background. Concealing coloration works only in appropriate surroundings and only when accompanied by proper behaviour, which is usually immobility. Great numbers of larval, pupal, and adult Lepidoptera are thus protected in their usual locations on plants. Leaf-eating larvae usually blend into leafy environments. Many caterpillars have stripes that simulate leaf veins. The saw-toothed elm caterpillar (Nerice) has a jagged outline resembling the edge of an elm leaf. A great many measuring worms, or inchworms, are notably twiglike, with long, slender, stiffly held bodies. Many other caterpillars, especially those of prominent moths (family Notodontidae), have irregular shapes that resemble twisted dead leaves. Likewise, many adult moths that rest during the day among leaves or on bark are cryptically coloured and patterned. Their behavioral mechanisms, such as aligning stripes on their wings with patterns of the tree bark, help them blend in with the backgrounds on which they rest.

Many larvae and adults are disruptively marked with bold contrasting patches or bands of colour that break up their outlines into two or more seemingly unrelated masses. Many adult moths and butterflies have coloration that serves to startle and thus momentarily delay an attacker. Moths with cryptic forewings and butterflies with cryptic wing undersides show only these surfaces when they are at rest. When moths such as the underwing moths (Catocala) are disturbed, they move the cryptic forewings to expose bright patches of colour on the upper surface of the hind wings. When butterflies such as the morphos, hairstreaks, and anglewings are disturbed, they take flight, exposing brightly coloured upper wing surfaces. Regardless of the method, the effect is startling to predators. When the animal lands, the bright surfaces are suddenly hidden, causing it to disappear into the background. A similar “startle effect” protects larvae that have prominent spots that look like large eyes.

Many species produce startling sounds. Hawkmoth caterpillars and the pupae of many gossamer-winged butterflies make squeaking or grating sounds when disturbed. The adult death’s head moth (Acherontia atropos) makes a loud chirping sound. Ageronia butterflies, when startled into flight, make a loud clicking sound by means of a structure on the wings. These sounds may have a startling and therefore delaying effect on a predator.

Certain butterflies and moths possess repellent or toxic substances that provide protection against predators. Sometimes these are secured directly from the plant on which the larva feeds, such as the toxic glycosides present in high concentrations within plants eaten by milkweed butterflies (family Nymphalidae) such as the monarch. More often, the toxin is produced by the insect itself and stored in the body, so that the predator must taste the insect to know it is toxic. The toxin often occurs in the blood—e.g., hydrogen cyanide in burnet moths. Other toxins may be in the gut or may be the product of special glands, which release the toxin at the time of an attack. Tiger moths (family Arctiidae) give off drops of repellent from glands on the prothorax. Many groups show reflex bleeding (autohemorrhization) from leg and body joints when disturbed. The larvae of swallowtail butterflies (Papilio) and tussock moths (family Lymantriidae) give off strong-smelling, volatile substances from extrusible scent organs (osmeteria). The caterpillars of many prominent moths spray formic acid from ventral prothoracic glands. Many larvae and some adults possess hollow barbed hairs that introduce toxins into potential predators, causing pain and swelling. Caterpillars of the slug moths (family Limacodidae), flannel moths (family Megalopygidae), Io moths, and some tussock moths are noted for this. A few adult moths (e.g., the garden, or great, tiger moth [Arctia caja]) inject toxins through sharp spines on their hind legs. The great majority of protected forms are aposematic; that is, they have markings, shapes, or behaviour that draw attention. They are thus easily recognized and remembered by predators, which, after trying to feed on only one or two individuals, will leave other similarly patterned individuals alone.

Not all warning mechanisms are visual. Inedible tiger moths make high-pitched grating sounds by means of timbal (drumlike) organs. These sounds are inaudible to humans but can be heard by bats, and they may function to warn the bat of the moth’s inedibility. This allows the moth to avoid being captured and tasted. Some authorities believe that these sounds also may interfere with the bats’ acoustic orientation system, preventing them from detecting the moth.

The protective advantage gained by a distasteful or dangerous insect that advertises that defense, either by aposematic coloration or acoustic warning, may also be utilized by similar harmless and edible insects. The evolution of such resemblances results in a phenomenon known as mimicry. (For background information on this phenomenon, see mimicry.) The distasteful insect, called the model, may even belong to an insect order completely different from that of the mimic. For example, members of various lepidopteran families mimic wasps, bees, or beetles. The clearwing moths are particularly effective mimics of certain stinging wasps, the resemblance being carried to details of the shape and coloration of the wings, abdomen, and legs.

The occurrence in a population of two or more distinct hereditary variants, or morphs, is known in many lepidopterans. Each morph may have a different adaptive value, linked with such physiological features as resistance to cold or to toxins in the environment. Striking variation in appearance may have great adaptive value by confusing predators, making it more difficult for them to learn the appearance of the prey. Mixed populations of both light and dark (melanic) individuals may survive better in habitats containing both light and dark backgrounds on which they rest during the day. This may also allow the entire population to survive if the environment changes either through normal succession of forest growth or because of man-made phenomena such as industrial pollution. In England investigations of the peppered moth Biston betularia have abundantly documented the evolution of “industrial” and “natural” melanism and have shown that major genetic population changes can take place very rapidly.

Striking polymorphisms occur in some mimetic species, notably the African swallowtail (Papilio dardanus). The occurrence of different species of inedible butterfly models in various geographic regions has been accompanied by the evolution of correspondingly different mimetic females of this single species of swallowtail. In North America the tiger swallowtail (P. glaucus) has mostly black females wherever it coexists with the distasteful pipevine swallowtail (Battus philenor), which is also black. However, where B. philenor does not occur, P. glaucus females tend to be all nonmimetic yellow forms like the males because, without the black models, black has no protective significance. Some very striking mimetic polymorphisms occur among Neotropical Heliconius butterflies and their various models and mimics.

Form and function

Within the limitations imposed by their almost entirely plant-eating habits, the lepidopterans show a great diversity of size, structure, and other characteristics. Furthermore, some members of the order have retained many primitive features, while others have evolved very advanced and specialized ones. This broad range, from primitive to advanced and from generalized to specialized, is evident not only in the adults but also in the larvae and pupae and to some degree in the eggs.

The adult

Nearly all external surfaces of the adult are covered with scales, which may be broad and flat or long and hairlike. Each of these is the outgrowth of a single epidermal cell. Similar scales occur in a few groups of other insects but never to the same extent.

Like most insects, adult lepidopterans have three distinct body segments, or tagmata—the head, the thorax, and the abdomen—each with special functions. The head bears the main sensory organs and those of feeding and ingestion. The thorax is chiefly concerned with locomotion. The abdomen contains the main organs of digestion, excretion, and reproduction and bears the external accessory reproductive structures.

Head

The head is relatively small and round or elliptical. With regard to its evolutionary development, it is derived from the first six primitive body segments (somites, or metameres), but these have become so coalesced that none of the primitive segmentation is evident. The antennae are prominent and multisegmented, with many microscopic receptors (sensilla) for detecting odours. In most moths the antennae are slender and tapering, although in some the surface area of the antennae is greatly enlarged by many side branches that make them comblike or featherlike. In skippers and butterflies the terminal part is expanded into a clublike shape, on which most of the sensilla are concentrated. In many families there is also a cluster of sensory bristles (the chaetosema) on each side of the head near the eye. On either side of the head is a large compound eye, sometimes consisting of thousands of units (ommatidia). Most moths have, in addition to the compound eyes, a pair of very small simple eyes (ocelli), which have limited light-sensing ability but do not form an image.

The compound eyes are very efficient for distinguishing motion but not for resolving clear images of distant objects. Diurnal species may be able to distinguish flower shapes, and courting individuals have been shown to respond to specific wing patterns. Colour perception, especially in the blue-violet end of the spectrum, is acute in some species. Moths are generally able to see by ultraviolet light, to which they are often strongly attracted at night.

The sense of smell is acute in some groups, especially in males with large antennal surfaces. The sense of taste, especially for sugars, is also acute. In moths and butterflies, taste receptors are located chiefly on the palpi and on the feet (tarsi).

Nearly all caterpillars have chewing mouthparts, but these are retained among adults of only the most primitive moths—the mandibulate moths of family Micropterigidae. Caterpillar mouthparts basically consist of an anterior flap (labrum), a pair of chewing jaws (mandibles), a pair of complex first maxillae, and a pair of similar second maxillae joined together behind the mouth to form a structure called the labium. Each of the first and second maxillae bears a jointed sensory appendage, or palpus. All these structures function together for chewing and manipulating solid foods. In the vast majority of adults the mandibles are either vestigial and nonfunctional or entirely absent. Parts of the first maxillae, however, are elongated to form the two halves of a tubular proboscis (haustellum) through which liquids may be sucked. The segmented palpi of the first and second maxillae are present and function as sensory organs. Not all adults have all these parts fully formed and functional. In numerous families the proboscis has become considerably reduced and even vestigial, resulting in adults that cannot feed. In advanced moths and in skippers and butterflies, the maxillary palpi are vestigial or lost, so that only the labial palpi remain functional.

Mouthpart reduction in adults appears to have occurred independently in a number of evolutionary lines. The result in these species has been to limit feeding to the larval stage and thus enable the adult stage to be dedicated only to reproduction and dispersal.

Thorax

The thorax consists of three segments, the prothorax, mesothorax, and metathorax, each derived from a primitive segment. The prothorax bears the first pair of legs and a pair of respiratory openings (spiracles). The much larger mesothorax bears the second pair of legs, a second pair of spiracles, and the pair of forewings. The metathorax bears the third pair of legs and the pair of hind wings. In many moths the metathorax bears a pair of complex auditory organs (tympana). In some species these organs serve as receptors of the high-frequency echolocation signals emitted by hunting bats, allowing the moths to initiate escape maneuvers. In other species the auditory organs are receptors of mate location calls. Sound signals are produced in some species by timbal organs and in others by a mechanical clicking of the wing base.

The wings begin development in the maturing larva as invaginations of the epidermis. As the pupa is formed, these fold out (evaginate) to lie externally and become large and flat. Within them, branching tubes (tracheae) carry an air supply and also mark the pathways along which will develop the tubular “veins” that support the fully formed wings. When the adult emerges from the pupa, the wings expand to full size. On the wing the scales lie in overlapping rows, like shingles. The usual scale is a flattened, rigid, air-filled sac attached by a peglike base. It is usually ribbed longitudinally and toothed terminally. Many males have special glandular scent scales (androconia) scattered or concentrated in patches (brands or stigmata) on the wings, sometimes forming expansible, hairlike tufts. These have a scent-distributing function that is essential in specialized courtship.

The scales and hairs of the adults, especially on the wings, are responsible for colours and patterns. There are many different pigments of several chemically diverse types such as melanins, uric acid derivatives, and flavones. In addition, the microstructure of hairs and scales refracts light in various ways so as to produce colours as well as metallic, pearly, iridescent, and white effects. Very fine parallel ridges in scales may produce an iridescence by acting as diffraction gratings. More common are the effects of very thin superimposed layers (laminae) in the walls of the scales, a colour phenomenon comparable to that produced by a thin film of oil on water. Many of the most striking effects are caused by combinations of pigmental and structural colours.

The forewings and hind wings on each side are coupled together in various ways. In primitive moths a fingerlike lobe on the forewing overlaps the base of the hind wing. In most moths a strong bristle or cluster of bristles (frenulum) near the base of the hind wing engages a catch (retinaculum) on the forewing. In some moths and in the skippers and butterflies, the frenulum mechanism has been lost, and the wings are coordinated by the friction of the overlapping areas. In the most primitive moths the forewings and hind wings are similar in size, shape, and veinage. In most moths and in the skippers and butterflies, the hind wings have become shorter and more rounded, with reduced veining except in the posterior wing section. The anterior, or leading (costal), edge of the forewing is thickened, with stronger veins, while the outer and posterior (anal) wing section margins are thinner and weaker. This accords with the function of the wings as airfoils having a stiff leading edge and a flexible trailing edge.

The wings are characteristically modified in various families. In many moths of the superfamily Tineoidea, both wings have become extremely narrow, with much-reduced venation. However, along the margins there are long, dense fringes of hairs that maintain the functional wing area. In the plume moths (family Pterophoridae) the wings are deeply cleft into two or three narrow plumes, and in the many-plumed moths (family Alucitidae) each wing is cleft into six plumes. In a number of moth families the females are wingless, although the males are fully winged (bagworm moths [family Psychidae], some tussock moths [family Lymantriidae]). In the aquatic snout moth (Acentropus) some females are wingless, while some females and males are winged. There are eight main wing veins, each with a characteristic pattern. These are usually designated according to the modified Comstock-Needham system. The names of the veins (with their symbols in parentheses) and the usual number of branches of each (subscript designations) are as follows, in progression from the costal margin of the wing to the anal margin:

Costa (C), along the anterior margin, no branches; Subcosta (Sc), usually with no branches, rarely with two (Sc1, Sc2);Radius (R), typically with five branches (R1–R5); Media (M), with three branches (M1–M3; four in some other insects);Cubitus (Cu), primitively with three branches (Cu1a, Cu1b, Cu2);Anal veins, First (1A), Second (2A), and Third (3A), all unbranched.

Abdomen and genitalia

The abdomen has 10 segments, although the posterior ones are indistinct. Each of the first eight segments bears a pair of spiracles. The first or second segments bear paired auditory organs in the snout moths and measuring worm moths. Segmental appendages are absent except for vestiges that may form parts of the genitalia. Various segments may bear special structures that produce and disperse pheromones. The genitalia of both sexes are often complex and bear characteristic spines, teeth, setae, and scale tufts. These structures are important in complex courtships and matings, preventing hybridization between males and females of different species.

In males a ringlike structure is the base of attachment for a number of dorsal structures and a pair of lateral clasping organs (valvae). In copulation a median tubular organ (the aedeagus) is extended through an eversible sheath (vesica) to inseminate the female. These structures are derived evolutionarily from parts of segments 8 and 10 and from vestiges of abdominal appendages.

The female genitalia show a number of different types of organization of the internal genital ducts and openings. These are considered so fundamental that the lepidopterans can be classified into suborders largely on the basis of these traits and their correlation with characteristics of the mouthparts, wings, and early developmental stages.

The internal reproductive systems of both sexes contain the organs typical of most insects. The testes of the male are paired in primitive lepidopterans but fused into a single organ in advanced forms. In both cases the sperm ducts are paired. As in other insects, the sperm pass from the testes down these paired ducts (vasa deferentia) for storage in sacs called seminal vesicles. Accessory glands, providing fluids that lengthen the life of the sperm, open into the vasa deferentia.

The female reproductive system consists of paired ovaries, paired accessory glands that provide the yolks and shells of the eggs, and a system of receptacles and ducts for receiving, conducting, and storing sperm. The individual oviducts join to form a common oviduct that leads to the vagina. In copulation the male deposits a sperm capsule (spermatophore) in a receptacle of the female. The spermatophore releases the sperm, which swim into the oviduct and thence to the seminal receptacle, where they are stored until egg laying. This may occur hours, days, or months after mating.

The egg

The egg is enclosed in a protective layer (the chorion), through which a system of tiny canals (micropyle) permits the entrance of sperm. In some groups the micropyle is at the side, whereas in others it is on the surface away from the substrate. The egg passes along the individual oviduct and through the common oviduct to the vagina. Here, just before it is laid, it comes into contact with a droplet of seminal fluid that has been stored in the female, and fertilization takes place. Most eggs are more or less spherical, but those of a few families are flat or long and tapered at the ends. Their surface may be strongly sculpted with pits, sharp projections, or raised ridges.

The larva, or caterpillar

Compared with the highly specialized adult, the larva is simple and primitive. Many of the primitive characteristics retained in the larva are important in the classification of the suborders, superfamilies, and families. The traits also aid in making speculations on relationships among these groups (taxa).

The head bears a pair of very short antennae and on each side a cluster of minute simple eyes (stemmata). A short liplike labrum is in front of the mouth. Behind the labrum are paired jaws (mandibles) that are short, broad, and powerful to allow consumption of large amounts of plant material. Next is a pair of small first maxillae, each with a segmented palp. Then, more or less connected with the maxillae, is the labium-hypopharynx, a complex structure with a pair of labial palps between which is located a tubular spinneret through which silk is extruded. Each of the three thoracic segments bears a pair of short segmented legs. The remaining 10 segments constitute the abdomen. Abdominal segments 3 through 6 and segment 10 bear a pair of fleshy appendages called prolegs, which may be homologous with the primitive segmental appendages. Each proleg has one or two curved rows of minute hooklets and an eversible soft end, the planta. The crochets on the prolegs allow the larva to hold onto surfaces. Body fluids forced into the proleg cause it to expand, extending the hooklets. After the proleg has been placed on the substrate, the fluids are retracted into the body and the elasticity of the cuticle causes the tiny hooks to retract, thus gripping the substrate. Prothorax and abdominal segments 1 to 8 have a pair of spiracles on them.

The larval epidermis bears on the head and each body segment a number of bristles known as primary setae. The position and number of setae are constant in each species and so are important in classification. Often there are many secondary setae, giving the caterpillar a hairy appearance. Larvae that live and feed as borers, burrowers, or miners are mostly plain. Those that live and feed in the open often show a great diversity of shape, colour, pattern, and ornamentation. Many have wartlike projections (verrucae) that may bear tufts of setae or spiny projections, or there may be prominent hornlike or spiny processes.

In some families the number of prolegs has been modified. In the measuring worms the prolegs of segments three, four, and five are missing, and in some owlet moths one or two pairs have been lost. In the puss moth caterpillars the last pair has evolved into a pair of long eversible whiplashes (stemmatopoda). The larvae of some leaf miners (family Gracillariidae) have lost some or all of the prolegs. Mandibulate moth larvae have eight pairs of abdominal legs, which are structured more like true thoracic legs than prolegs.

Internally, the larva is relatively simple, with the very large digestive tract being the most prominent organ. The paired silk glands are often very large, extending far back into the abdomen. The ovaries and testes, which begin to develop during embryonic life, continue to develop in the larva, as do the wings. Many special glands secrete repellent or toxic substances, which may circulate in the blood or be extruded from special openings as a means of defense.

Larval vision can detect little more than differences between light and darkness. Taste is acute, with highly developed sensory receptors in the antennae and palpi. Food discrimination is keen, and many larvae will starve rather than eat abnormal food plants. The sense of touch functions via setae widely distributed over the outer surface. Some of these appear to react to sound waves of low pitch, well within the limits of human hearing.

The pupa, or chrysalis

Lepidopteran pupae show the same sort of evolutionary gradation from primitive to advanced as do larvae and adults. In the primitive mandibulate moths and sparkling archaic sun moths (family Eriocraniidae), the pupa has free and movable appendages and functional mandibles. In some less-primitive groups the pupa retains the ability to move some appendages. In the higher moths and the butterflies, all appendages are tightly fastened to the body wall. Called obtect pupae, these are immobile and able to wriggle only one or two abdominal segments. In a few groups the pupa has special stridulating rasps for sound production. Nearly all of the external structures of the adult can be seen on the pupa. The wings are prominent, folded down flat along the ventral surface, with the proboscis halves, the legs, and the antennae between them. At the posterior end is a spiny pad or spike, the cremaster, which in many groups attaches the pupa to silk fibres spun by the larva.

Except for a very slight respiratory exchange and a little water loss, the pupa is physiologically self-sufficient. Within it most of the cells and tissues of the larva undergo considerable histolysis (breakdown) as the adult structures are built up from the existing rudiments. Some structures begin developing as far back as the first larval stages.

Growth, molting, and metamorphosis

As in other insects, growth and its structural changes are controlled by an interacting set of hormones. These are chiefly secreted by the corpora allata and other parts of the brain and by paired prothoracic glands. The prothoracic gland hormone is necessary for larval molting (ecdysis), metamorphosis to the pupa, and formation of adult characteristics. On the other hand, a hormone secreted by the corpora allata inhibits metamorphosis until late larval development. A hormone secreted by cells in the pupal brain stimulates the prothoracic glands and thereby brings about differentiation of the adult and the end of the obligatory resting stage (diapause) of the pupa.

Evolution and paleontology

The Lepidoptera belong to an important group of insect orders called the panorpoid complex. This ancestral stem began in the Permian Period (290 299 million to 248 251 million years ago) and split into a number of branches, from which evolved the modern scorpionflies (order Mecoptera); dobsonflies, alderflies, and lacewings (see neuropteran); true flies (order Diptera); caddisflies (order Trichoptera); and the lepidopterans. The nearest living relatives of the Lepidoptera are the caddisflies, and in fact the very primitive mandibulate moths (family Micropterigidae) have been grouped with the caddisflies by some systematists. As the Lepidoptera developed into a distinctive group, certain major trends began to be emphasized. The adult mandibles disappeared and were replaced by the proboscis formed from the galeae of the maxillae, which broadened the food base exploitable by the species to include not only sap and other plant juices but nectar and fruit juices as the flowering plants evolved. Various groups, however, independently lost the proboscis and concentrated on the larval stage as the sole source of nutrition. The characteristic vein pattern evolved, with the long, veinless discal cell in each wing, as well as an emphasis on the strength of the forewing and a de-emphasis of the hind wing, forming a particular pattern of aerodynamic efficiency. Quite a few groups of small moths, settling into restricted ecological niches where strong flight is not a necessity, evolved a great deal of wing reduction.

Concurrent with changes in the adults, the larvae were evolving a multiplicity of different ways of feeding on the evolving seed plants. As both larval and adult differentiations accumulated, it became more and more necessary that the pupal stage be a passive, resting phase to allow time for the metamorphosis of larva into adult.

Very few fossil Lepidoptera are known. The earliest, dating from the beginning of the Paleogene Period, which began 65.5 million years ago, are small moths related to the superfamilies Eriocranioidea and Tineoidea. However, given their characteristics, the order must have existed long before then. The closely related caddisflies are known from the Jurassic Period (206 200 million to 144 146 million years ago). The best-known fossil butterflies are relatively recent, found in the shales of the Florissant Formation, a deposit laid down in Colorado at the Eocene-Oligocene boundary some 34 million years ago. Some of those are undoubtedly brush-footed butterflies (family Nymphalidae), which are very similar to modern genera.

Classification

Distinguishing taxonomic features

The chief characteristics used in the ordinal, subordinal, and family-level classification of the Lepidoptera are: the mouthparts, ranging from mandibulate to haustellate (with a proboscis) or degenerate; the venation and shape of the wings, homoneurous (the venation of the forewings and hind wings alike) or heteroneurous (forewings and hind wings different), aculeate (more or less covered with specialized bristles called microsetae) or nonaculeate, and type of coupling (jugum or frenulum); the anatomy of the female reproductive system; the exarate or obtect condition of the pupa; and the larval structure and pattern of the primary setae.

The venation of the wings is perhaps the most important single criterion for establishing both differences and relationships in the classification. However, venation patterns must be considered in terms of the evolution of these patterns from primitive to advanced conditions within individual phyletic lines. The most primitive groups tend to have the maximum number of veins and branches in each wing. More advanced groups lack some veins or branches in the forewing and have the hind-wing venation considerably reduced.

The structures of the genitalia are extremely important in classification, often serving better than any other characteristics to distinguish the species. The various parts have been homologized with each other, thus enabling comparative studies of families, genera, and species.

The female genitalia exhibit a number of different patterns of the internal ducts and the openings. These vary from a condition in which there are no special genital openings, with insemination and egg laying (oviposition) taking place through a single aperture shared with the excretory system, to one in which there are two specialized openings, one for insemination and one for oviposition, both of which are distinct from the anus.

Annotated classification

The classification presented follows Niels P. Kristensen (1999). Classification systems vary in the way they categorize taxonomic groups. Traditional taxonomic schemes gather superfamilies and families into suborders; more recent systematic schemes have abandoned suborders in favour of less rigidly organized clades. In the scheme followed below, neither suborders nor clades are indicated; instead, only superfamilies and families are shown. Of the 46 lepidopteran superfamilies, 24 are presented below, representing all the commonly encountered families of moths, butterflies, and skippers. Superfamilies and families not presented below include all moths that are rarely seen.

Order LepidopteraInsects with complete metamorphosis; the wings covered with flat scales; larval mouthparts mandibulate; adult mouthparts mandibulate to haustellate, sometimes vestigial; wings with venation ranging from primitive (complex) to considerably reduced, especially in the hind wings; pupae exarate to obtect; almost all phytophagous (herbivorous).Superfamily NoctuoideaAlmost 52,000 species in 8 families; adults with a pair of complex tympanic organs on metathorax.Family Noctuidae (owlet moths)More than 35,000 species worldwide; abundant everywhere with a great diversity of size, colour, and habit; sizes range from a wingspan of 10 mm (0.4 inches; Hypenodes) to 275 mm (Thysania); many species brightly coloured, but the majority are plain and cryptic; nearly every type of plant material is exploited as food: foliage, flowers, fruits, stems, rootstocks, and woodland litter; a few genera (e.g., Cosmia) cannibalistic; family includes the armyworms and cutworms, among the most injurious of all moth larvae to human interests.Family Arctiidae (tiger moths)Approximately 10,000 species worldwide; adults usually brightly coloured, especially in tropics; aposematic coloration, with protective and toxic secretions; timbal organ for making very high-pitched sounds located on each side of the metathorax; larvae often with much secondary hair; several other families, chiefly tropical, are closely related and also have timbal organs.Family Lymantriidae (tussock moths)More than 2,500 species worldwide, but mainly in Old World tropics; adult females heavy-bodied, sometimes wingless; many larvae with prominent tussocks and pencils of hair, poison spines, and protrusions that emit repellent odours (osmeteria); some—e.g., the gypsy moth (Lymantria dispar) and the nun moth (L. monacha)—are very destructive to forests; closely related Thaumetopoeidae of the Old World are best known for the gregarious habits of the larvae (processionary caterpillars), which move in orderly columns.Family Notodontidae (prominent moths)More than 2,800 species worldwide, except New Zealand and Pacific islands; adult coloration mostly dull, cryptic; larvae very diverse with cryptic, disruptive, or aposematic coloration; some protected by toxic secretions; posterior prolegs often modified or aborted; chiefly foliage eaters.Superfamily PyraloideaApproximately 17,800 species in 2 families worldwide; most with a pair of tympanal organs on the first abdominal segment; adults usually slender-bodied with long legs; many with narrow forewings and broad, often folded, hind wings.Family Pyralidae (pyralid, or snout, moths)Approximately 6,130 species of small moths, mostly plain, often abundant, with many important pest species; differentiated from other families by wing venation; small subfamily Nymphulinae has aquatic larvae with tracheal gills for living in still or running fresh water; larvae of subfamily Pyralinae are mostly scavengers, as are those of the Galleriinae, many of which live in bee or wasp nests; larvae of the large subfamily Phycitinae have very diverse habits, including predation on scale insects.Family Crambidae (webworms)Approximately 11,600 species worldwide; small, often abundant moths, many larvae producing silk webbing in feeding sites; subfamily Crambinae contains almost 1,900 species, larvae feeding mainly on roots, grasses, or mosses on the ground or boring into stems of grasses, sedges, or rushes; subfamily Pyraustinae contains more than 7,400 species, feeding mainly on stems and fruits of various plants; many Pyraustinae species are considered pests, but some have been used in management of aquatic weeds.Superfamily GeometroideaAlmost 22,000 species; adults with abdominal tympana; some authorities classify each of the 3 major families as a separate superfamily.Family Geometridae (measuring worm, or inchworm, moths)Approximately 21,000 species, abundant worldwide; adults and larvae commonly very cryptic, resembling bark, dead leaves, and twigs; larvae (“inchworms”) long, slender, lacking most posterior abdominal prolegs, crawling with characteristic looping gait; pupation usually in the soil, with no cocoon; many species destructive to foliage—e.g., cankerworms (Alsophila and Paleacrita) and the winter moth (Operophtera brumata). Family Uraniidae (swallowtail moths)Approximately 700 chiefly tropical species; some adults are large, brilliantly iridescent diurnal moths; the Asian Epicopeia (family Epicopeiidae) mimic swallowtail butterflies.Superfamily DrepanoideaApproximately 700 species worldwide in 2 families.Family Drepanidae (hooktip moths)Approximately 650 species worldwide, chiefly Indo-Australian; many of the adults have the forewing apexes strongly hooked; larvae usually lack last pair of prolegs; subfamilies Thyatirinae and Epibleminae sometimes classified as families.Family Epicopeiidae (epicopeiid moths)25 species in Arctic and tropical Asia; colourful day-flying moths that often mimic butterflies and other colourful moths such as the Arctiidae; larvae feed on foliage of woody plants.Superfamily GelechioideaMore than 16,000 species worldwide; adults mostly larger and broader winged than Tineoidea; larvae seldom leaf miners; pupae relatively immobile.Family Gelechiidae (twirler moths)More than 4,500 species of small to minute moths, worldwide in distribution; larvae diverse, eating leaves, stems, fruit, and tubers, including a few gall makers; serious economic pests include the pink bollworm of cotton and the Angoumois grain moth (Sitotroga cerealella).Family Cosmopterigidae (cosmopterigid moths)More than 1,600 species of small moths, worldwide in distribution; many adults are very narrow-winged with bright, often metallic markings; in addition to leaf miners, rollers, and tiers, larvae include stem, fruit, and seed borers as well as scavengers.Family Coleophoridae (casebearer moths)Approximately 1,400 species, mainly Holarctic in distribution; small, very narrow-winged moths; larvae mostly mine leaves or feed on seeds; many larvae construct portable cases with distinctive shapes; some are pests of fruit trees.Family Oecophoridae (oecophorid moths)More than 3,100 small species worldwide; adults tend to be flat-bodied and somewhat broader-winged than related groups; related families: Elachistidae, Xylorictidae.Superfamily Papilionoidea (butterflies)14,000 species, all families worldwide; adults with clubbed but not hooked antennae; flight slower than moths and not darting; larvae lacking “neck”; general evolutionary trend has been from a primitive condition of fully developed forelegs in both sexes (Pieridae, Papilionidae) to one in which they are greatly reduced and useless for walking (Nymphalidae); intermediate conditions occur in Lycaenidae; pupae (“chrysalides”) often brightly coloured and irregularly shaped, not enclosed in cocoons.Family Lycaenidae (blues, coppers, hairstreaks, and metalmarks)6,000 small and diverse species; many iridescent blue, green, or metallic orange; some adults have thin tails on the trailing margin of the hind wing; larvae somewhat sluglike, hairy, many secreting honeydew and having mutualistic relationships with ants; some prey on the ant brood or on other insects; many pupae have stridulatory structures on the abdomen.Family Nymphalidae (brush-footed butterflies)Approximately 6,000 species, often split into several families that here are considered subfamilies; Nymphalinae is the main subfamily, with many familiar species such as the fritillaries, admirals, checkerspots, and anglewings; many tropical species brilliantly iridescent; Satyrinae contains the familiar wood nymphs, meadow browns, and heaths, usually with eyespots on the wings; larvae distinctively pointed at the rear; spin crude cocoons; the Libytheinae (snout butterflies) are so named because of their long protruding palps; the very large Brassolinae and iridescent Morphinae are Neotropical, as are the highly distasteful, aposematic Heliconiinae and Ithomiinae that, with the worldwide Danainae, are models in many mimicry complexes; most of the pantropical Acraeinae are also highly protected and aposematic models; some nymphalids, such as the monarch butterfly, are migratory.Family Pieridae (white, orange-tip, and sulfur butterflies)Approximately 1,000 small to medium-size species; no native species are found in New Zealand; mostly white, yellow, or orange, often with dark tips on wings; pupae usually with a frontal horn on head; many species noted for mass migrations.Family Papilionidae (swallowtails and parnassians)600 medium to large species, often brightly coloured; many have the “swallowtails” on the hind wings; some females of the Indo-Australian birdwings (Troides) are the largest butterflies; larvae often curiously patterned, with eversible, protective scent organs (osmeteria); adults of many are highly distasteful and much mimicked; parnassians sometimes placed in a separate family, Parnassiidae.Superfamily Tortricoidea6,100 species in 1 family; adults with fairly broad, short-fringed wings that seldom span more than 2.5 cm (1 inch); most have cryptic coloration and patterns; larvae mostly leaf folders and rollers, but many bore in fruits, seeds, and soft stems.Family Tortricidae (leaf roller moths)Approximately 6,100 species worldwide; family large and diverse; most larvae are stem borers or leaf rollers or feed in leaf litter; larvae of the green leaf roller of Europe (Tortrix viridana) defoliate oak forests; the spruce budworm (Choristoneura fumiferana) is the worst forest pest of North America. Superfamily TineoideaMore than 4,000 species worldwide; a large group of families of mostly small moths of diverse habits; all have some primitive venation features and life cycles; wings narrow to very narrow.Family Tineidae (clothes moths and other tineid moths)Approximately 3,000 species worldwide; small narrow-winged moths with rough, hairy heads; larvae often casemakers, feeding on debris and fungi; clothes moths (Tineola, Tinea, Trichophaga) often serious household pests; related family: Acrolophidae (burrowing sod webworms).Family Psychidae (bagworms)Almost 1,000 species worldwide; larvae live and pupate in often elaborate cases; adult males with broad, thin scaled wings; females wingless, often greatly degenerate and never leaving larval cases.Superfamily GracillarioideaApproximately 2,300 species worldwide; small moths; larvae are mainly leaf miners or stem borersFamilies Gracillariidae and DouglasiidaeApproximately 2,000 species worldwide whose larvae have degenerative legs and mandibles; adults with narrow, long-fringed wings often with metallic markings; larvae mostly leaf miners or stem borers, sometimes greatly flattened.Superfamily Hesperioidea 3,500 species worldwide in 1 family; similar to true butterflies, distinguished from moths by diurnal habits, clubbed antennae, a functional proboscis, and lack of ocelli; adults are fast-flying, with short, usually pointed forewings, broad heads, and antennae usually hooked beyond the club; larvae usually have a pronounced necklike collar.Family Hesperiidae (skippers)3,500 species worldwide; adults range from small and drab to large and iridescent, especially in the tropics; larvae mostly live concealed in individual leaf nests or in webs among grasses, forming flimsy cocoons. Superfamily BombycoideaApproximately 3,400 species; adults large to very large; male antennae comblike in form.Family Bombycidae (silkworm moths)350 species worldwide except Europe; most common in Asian and New World tropics; includes the domesticated silkworm (Bombyx mori); related family: Eupterotidae.Family Saturniidae (giant silkworm moths)1,480 large species worldwide; larvae very large and fleshy, often with brightly coloured knobs and spines; most species spin firm cocoons of brown, green, or silvery silk; adults often with bright colours and striking wing shapes and patterns; males with broadly branched antennae, wingspan up to 275 mm (10.8 inches) in hercules moth (Coscinoscera hercules) of Australia and New Guinea, though every continent has some well-known species—e.g., the North American cecropia (Hyalophora cecropia), Io (Automeris io), polyphemus (Antheraea polyphemus), and luna (Actias luna); European giant peacock or emperor (Saturnia pavonia); and Indo-Australian atlas moth (Attacus atlas). Subfamily Citheroniinae (regal moths) are medium to very large New World species of heavy-bodied moths, often brightly coloured; larvae often with long spines—e.g., the North American Citheronia (hickory and pine horned devils); pupation in the ground, with no cocoon; related families: Brahmaeidae (African and Indo-Australian), Mimallonidae (New World).Family Sphingidae (hawk, or sphinx, moths)1,200 medium-size to large species worldwide; adults powerful fliers with long, narrow forewings, thick antennae, and generally strong proboscises; feed by hovering before flowers and drawing nectar through the extended proboscis; some are diurnal, most are active at twilight (crepuscular); larvae large, fleshy, and mostly smooth; most have a single long posterior horn; typically rest with head and thorax reared, fancifully like the Sphinx at GazaGiza, Egypt, in profile.Superfamily LasiocampoideaApproximately 1,600 species worldwide.Family Lasiocampidae (tent caterpillar and lappet moths)1,500 species worldwide; larvae usually hairy and brightly coloured, some living gregariously in silk nests; adults medium-size, stout-bodied, short-winged, and exceptionally hairy.Superfamily ZygaenoideaApproximately 2,500 species worldwide; wings broad to very broad with short fringes; some primitive adult, pupal, and larval features; pupae quite mobile.Family Limacodidae, or Eucleidae (slug caterpillar moths)More than 1,000 species worldwide; larvae broad and flat, with reduced prolegs; move glidingly with head hidden beneath prothorax; many with toxic, irritant setae; adults with heavy hairy bodies and vestigial proboscises. Family Megalopygidae (flannel moths)240 species in Central and South America; larvae similar to those of Limacodidae, but with normal prolegs and traces of additional ones; setae very toxic and nettling. Family Zygaenidae (burnet and forester moths)More than 1,000 species, mainly in subtropical and tropical Asia and Palearctic regions; adults usually diurnal flower visitors, with bright colours and strong proboscises; strongly protected by high concentrations of hydrogen cyanide in blood; larvae are leaf skeletonizers; related families: Aididae and Chalcosiidae (Old World tropics); Pyromorphidae and Dalceridae (New World).Family Epipyropidae (parasitic moths)40 chiefly Asian species; larvae live as external parasites on plant hoppers; related family: Cyclotornidae (Australian; larvae live similarly when young, then move to ants’ nests).Superfamily YponomeutoideaMore than 1,500 species worldwide; a limited and not very distinctive superfamily; larvae possess distinctive primary setation.Family Yponomeutidae (ermine moths)Almost 600 species worldwide; adults brightly coloured, especially in the tropics; pupae of some make lacework cocoons; related families: Plutellidae, Glyphipterygidae, Heliodinidae, Scythrididae.Superfamily SesioideaApproximately 1,200 species worldwide; most sesioid moths are diurnal with many aposematic adults.Family Sesiidae (clearwing, or wasp, moths)More than 1,000 species worldwide; adults diurnal flower visitors; often brightly coloured with yellow, orange, or scarlet, the wings usually mostly transparent, often very striking mimics of wasps; larvae often are stem, twig, and root borers, often injurious to fruit trees.Family Castniidae (castniid moths)Approximately 130 species in Central and South America; medium-size to large diurnal species of the New World and Indo-Australian tropics; adults powerful fliers, heavy-bodied and broad-winged; clubbed antennae, bright colours; often mimic other butterflies and diurnal moths; larvae are often stem borers.Superfamily CossoideaApproximately 700 species described; adults range from small to large, usually robust moths; males often with bipectinate antennae; larvae mainly stem or wood borers.Family Cossidae (carpenterworm and goat moths)Almost 700 species described worldwide; medium-size to large moths; adults are the heaviest adult lepidopterans, with wingspans up to 25 cm (10 inches); larvae are wood borers, penetrating even solid heartwoods; may require at least 2 years to mature; some very destructive, including the coffee borer (Zeuzera coffeae) and leopard moth (Z. pyrina).Superfamily PterophoroideaAlmost 1,000 species in 1 family.Family Pterophoridae (plume moths)Almost 1,000 mainly tropical species; adults with very long, slender legs and bodies, the wings usually deeply cleft into plumes; larvae spin webs on and eat the leaves of various plants or bore into seeds, seedpods, roots, or stems.Superfamily AlucitoideaAlmost 150 species worldwide; this superfamily and the related Pterophoroidea are the only families with deeply lobed wings.Family Alucitidae (many-plumed moths)130 species worldwide; each wing is very deeply cleft into 6 or more narrow plumelike divisions.Superfamily NepticuloideaApproximately 900 species worldwide; females with one genital opening and a soft ovipositor. Family Nepticulidae (midget moths)800 species worldwide of very small to minute moths; antennae with broad “eyecaps” at the base; larvae mostly leaf and bark miners, a few gall makers. Family OpostegidaeApproximately 100 worldwide species of small moths with narrow long-fringed wings; larvae leaf, stem, or bark miners.Superfamily TischerioideaApproximately 80 species in a single family.Family Tischeriidae (trumpet leaf miner moths)Approximately 80 species predominantly in North America; not found in Australia or the rest of Oceania.Superfamily IncurvarioideaMore than 500 species; all females with an extensible, piercing ovipositor for inserting eggs into plant tissue.Family Incurvariidae (fairy, or leafcutter, moths)Approximately 100 species worldwide; many are small brilliantly coloured diurnal flower visitors; male antennae often several times as long as forewings; mutualistic relationships of the yucca moths (Prodoxinae) with their food plants are notable as an example of coevolution; family sometimes split into families Incurvariidae, Adelidae, and Prodoxidae; related family: Heliozelidae (shield bearers).Superfamily Hepialoidea520 species; females with two genital openings; adult mouthparts reduced and nonfunctional, antennae very short.Family Hepialidae (swifts, or ghost moths)Almost 500 species found worldwide but chiefly in Australia and New Zealand; medium-size to very large moths, some brilliantly coloured; larvae mostly bore in turf or wood; related families, less-known: Prototheoridae (Africa and Australia), Palaeosetidae (Australia), Anomosetidae (Australia).Superfamily MicropterigoideaThe most primitive lepidopterans; females with no special genital opening; larvae, pupae, and adults with mandibulate mouthparts.Family Micropterigidae (mandibulate moths)120 small species, a few found in the Northern Hemisphere, more in Australia and New Zealand; adults eat pollen; larvae eat mosses and liverworts.Superfamily Eriocranioidea24 species in 1 family; females with one genital opening, a long cloaca, and a piercing ovipositor; adults with a short proboscis; pupae with functional mandibles.Family Eriocraniidae (sparkling archaic sun moths)24 species with a Holarctic distribution; often brilliantly coloured; adults feed on nectar; related families: Mnesarchiidae (New Zealand), Mesopseustidae (India and Taiwan).